New Freshwater Snails of the Genus Pyrgulopsis (Rissooidea: Hydrobiidae) from California

THE VELIGER CMS, Inc., 1995 The Veliger 38(4):343-373 (October 2, 1995)

ROBERT HERSHLER

Department of Invertebrate Zoology (Mollusks), National Museum of Natural History, Smithsonian Institution, Washington, DC. 20560, USA

Abstract. Seven new species of Recent springsnails belonging to the large genus Pyrgulopsis are described from California. Pyrgulopsis diablensis sp. nov., known from a single site in the San Joaquin Valley, P. longae sp. nov., known from a single site in the Great Basin (Lahontan system), and P. taylori sp. nov., narrowly endemic in one south-central coastal drainage, are related to a group of previously known western species also having terminal and penial glands on the penis. Pyrgulopsis eremica sp. nov., from the Great Basin and other interior drainages in northeast California, and P. greggi sp. nov., narrowly endemic in the Upper Kern River basin, differ from all other described congeners in lacking penial glands, and are considered to be derived from a group of western species having a small distal lobe and weakly developed terminal gland. Pyrgulopsis gibba sp. nov., known from a few sites in extreme northeastern California (Great Basin), has a unique complement of penial ornament consisting of terminal gland, Dg3, and ventral gland. Pyrgulopsis ventricosa sp. nov., narrowly endemic in the Clear Lake basin, is related to two previously described California species also having a full complement of glands on the penis (Pg, Tg, Dgl-3) and an enlarged bursa copulatrix.

INTRODUCTION in the literature (Hershler & Sada, 1987; Hershler, 1989; Hershler & Pratt, 1990). The author recently reviewed the taxa belonging to Pyr- To gain a better understanding of the California Pyr- gulopsis Call & Pilsbry, 1886, the second largest genus (65 gulopsis, the author recently collected material of this genus Recent species) of freshwater gastropods in North America from various parts of the state. Represented among these (Hershler, 1994). The focus of that paper was on previ- collections were seven new species, which are described ously described forms, although it was recognized that below. While these species are discussed within the context perhaps an equal number of undescribed species await of the preliminary phylogenetic hypothesis proposed for attention in the western United States. While such nov- the genus by Hershler (1994), a revised analysis incor- elties abound throughout many of the western states, Cal- porating these new data is deferred pending description of ifornia, which has only 10-11 currently recognized species the many other new species known from the West. (one may now be extinct in the state), may harbor an especially large group, given that this huge state is well- watered and has an extremely complex hydrographic his- MATERIALS AND METHODS tory owing to tectonism and associated climatic factors (Minckley et al., 1986). Published taxonomic studies on Institutional acronyms are USNM (National Museum of hydrobiid snails and other small freshwater gastropods Natural History, Smithsonian Institution), and WBM from California are few (Taylor, 1981), and collecting (personal collection of Walter B. Miller, now housed [un- efforts have been largely concentrated in only four regions: catalogued] at Santa Barbara Museum of Natural His- Great Basin and Klamath system to the northeast, lower tory). Anatomical study was of alcohol-preserved snails Sacramento system in the San Francisco environs, Death that had been relaxed with menthol crystals and fixed in Valley region, and south-coastal and Salton Sea drainage dilute (about 4%) formalin. Methods of anatomical study, to the southwest. Of these, only the Death Valley region terminology, and characters are those of Hershler (1994). hydrobiids have been thoroughly surveyed and documented Anatomical illustrations were prepared from camera lu- Page 344 The Veliger, Vol. 38, No. 4

Table 1 Shell parameters for new species of Pyrgulopsis. fi = mean, s = standard deviation, SH = shell height, SW = shell width, HBW = height of body whorl, WBW = width of body whorl, AH = aperture height, AW = aperture width, W = whorl expansion rate, D = distance of generating curve from coiling axis, T = translation rate, AS = aperture shape. Measurements are in mm.

WH SH SW HBW WBW AH AW W D T AS P. diablensis USNM 883791 M 4.90 3.76 236 273 2.07 1.70 1.28 1.80 0.54 5.64 1.34 n = 17 s 0.22 0.30 0.22 0.18 0.12 0.11 0.09 0.15 047 1.14 0.07 P. eremica USNM 858264 M 4.03 1.90 1.28 1.42 1.15 0.82 0.68 2.28 036 5.29 1.20 n = 15 s 0.13 0.16 0.11 0.12 0.09 0.82 0.05 032 045 0.64 0.06 USNM 858265 M 410 2.17 1.43 1.62 1.28 0.95 0.77 2.12 0.61 5.31 1.24 n = 15 s 0.16 018 0.09 0.11 0.07 0.07 0.04 0.26 0.04 0.92 046 USNM 858266 M 4.48 2.66 1.73 1.95 1.54 1.20 0.94 2.12 0.57 5.72 1.27 n = 11 s 0.21 0.22 0.11 0.15 0.11 048 0.06 0.27 043 0.58 0.04 USNM 858267 M 408 1.89 126 1.41 1.13 0.81 0.66 229 036 5.49 1.23 n = 12 S 020 0.12 007 0.07 045 0.05 0.04 0.42 0.03 0.47 0.06 USNM 858271 M 4.18 2.67 1.99 212 1.70 1.33 1.03 2.43 0.55 4.09 1.30 n = 15 s 0.22 0.17 0.10 0.10 048 0.07 045 0.42 0.04 0.45 0.04 USNM 873138 M 4.14 2.45 1.72 1.86 1.47 1.13 0.93 2.28 0.60 4.63 1.21 n = 14 s 0.16 0.15 0.11 0.11 0.08 0.09 0.06 0.23 0.04 0.32 0.05 P. gibba USNM 858275 M 4.25 293 1.99 226 1.70 1.37 1.13 2.04 0.60 5.10 1.21 n = 10 s 0.20 0.16 0.12 0.13 0.10 0.09 048 0.13 0.05 0.70 0.03 USNM 858273 M 4.33 3.34 2.28 234 1.94 1.60 1.30 2.09 038 5.02 1.24 n = 10 s 0.17 0.13 0.10 0.11 048 0.06 0.07 0.27 045 0.38 0.05 P. greggi USNM 874139 M 4.38 231 1.47 1.68 1.27 0.99 082 2.06 0.60 5.24 1.21 n = 15 s 0.27 0.16 0.09 0.11 0.11 0.07 0.04 0.45 003 0.67 0.04 USNM 874140 M 4.21 1.74 1.21 132 1.06 0.77 0.75 232 0.61 4.66 1.17 n = 13 s 033 0.10 007 0.07 0.05 0.77 0.72 0.60 0.06 035 046 P. longae USNM 858262 M 4.23 2.76 1.79 2.03 1.53 1.24 0.97 2.30 037 5.25 1.30 n = 15 s 0.24 018 0.15 0.15 0.11 0.10 0.06 039 0.05 0.94 0.05 P. taylori USNM 883792 M 463 2.28 1.28 1.59 1.11 1.05 073 1.71 036 6.00 1.44 n = 17 s 037 0.19 012 0.13 008 0.07 047 0.14 047 1.25 0.08 USNM 883789 M 463 2.80 1.67 2.03 1.41 1.30 0.90 1.85 034 5.00 1.46 n = 16 s 037 0.24 0.15 0.13 0.10 0.10 0.09 0.18 0.05 0.84 0.10 P. ventricosa USNM 883790 M 4.14 237 1.66 1.83 1.45 1.18 0.91 2.31 039 5.04 1.29 rc = 16 s 016 0.12 012 0.10 0.09 0.09 0.04 038 0.05 0.72 0.08 cida drawings. Methods of shell measurements are those Diagnosis: The genus was recently diagnosed by Hershler of Hershler (1989); data are presented in Table 1. (1994).

SYSTEMATICS Pyrgulopsis diablensis Hershler, sp. nov. Family HYDROBIIDAE (Figures 1-3, 5A) Subfamily NYMPHOPHILINAE Etymology: Referring to occurrence of this species in the Pyrgwbffif Call & Pilsbry, 1886 Diablo Range of central California. Type Species: Pyrgula nevadensis Stearns, 1883; original des- Diagnosis: Shell ovate- to narrow-conic, medium-sized, ignation. umbilicate. Penial filament medium length; lobe short. R. Hershler, 1995 Page 345

Figure 1 Scanning electron micrographs of shells of P. dtablensis Hershler, sp. nov., USNM 883791. F, Protoconch, har = 150 urn. Shell "A" is 3.8 mm tall; other shells primed to same scale.

Penial ornament a short penial gland and small terminal usually slightly separated from body whorl. Inner lip com- gland. plete, thin, without columellar shelf. Outer lip thin, slightly prosoeline. Umbilicus rimate to perforate, Feriostracum Description: Shell (Figures 1, 5A) ovate- to narrow-conic; light brown. height 3.1-4,3 mm; whorls 4.25-5,0. Protoconch (Figure Operculum (Figure 2A-C) ovate, amber with brown to IF) slightly less than 1.5 whorls, early portion strongly red attachment scar region; nucleus eccentric; dorsal sur- wrinkled. Teleoconch whorls convex, sometimes strongly face frilled. Attachment scar thickened, often strongly so, shouldered; sculpture of strong growth lines. Aperture ovate, all around. Callus large, well developed. Page 346 The Veliger, Vol. 38, No. 4

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Figure 3 Genital morphology of P. diablensis Hershler, sp. nov., USNM 883791. A. Distal female genitalia (left side). Bar = 0.5 mm. B. Penis (dorsal aspect, left; ventral aspect, right). Bar = 0.5 mm. Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled oviduct, Pg = penial gland, Sr = seminal receptacle, Tg = terminal gland, Vc = ventral channel.

Buccal mass large; radular sac protruding behind buccal idial axis. Kidney with medium pallial bulge; renal gland mass as short coil. Radula ribbon elongate, about 0.89 x longitudinal-slightly oblique; kidney opening white. Rec- 0.11 mm, with about 60 rows of teeth. Central radular tum straight, broadly overlapping genital ducts. Hypo- tooth (Figure 2D) trapezoidal, with medium to highly branchial gland medium thickness; overlapping rectum, indented dorsal edge; lateral cusps 4-5; central cusp slight- pallial gonoducts, narrow section of posterior pallial roof ly pointed, considerably broader and longer than laterals; adjacent to rectum. basal cusps 1, medium-sized, with medium dorsal support. Distal female genitalia shown in Figure 3A. Ovary 1.5 Basal process medium width; basal sockets deep. Lateral whorls, slightly overlapping posterior stomach chamber. margins slightly thickened, with weakly developed neck. Pallial albumen gland short. Capsule gland considerably Lateral tooth (Figure 2E) formula 3-1-3(4); neck weakly shorter than albumen gland. Genital aperture a broad slit, developed; basal tongue well developed; outer wing about with short vestibule. Coiled oviduct circular-oblique, kinked 140% of cutting edge length. Marginal teeth (Figure 2F, at mid-point, augmented by proximal twist or small coil. G) with about 15-19 (inner) and 16-23 (outer) cusps; Oviduct and bursal duct join slightly behind pallial wall. cusps extending onto outer edge of teeth. Stomach caecum Bursa copulatrix broadly ovate-triangular, about one-fourth small. length of albumen gland, and about two-thirds width of Cephalic tentacles pale, but with light grey internal gland, with three-fourths or more of length posterior to pigment proximally. Snout, foot grey to black. Opercular gland. Bursa duct narrow-medium width, about 150% of lobe black along inner edge, sides; outer edge pale to black. length of bursa copulatrix, shallowly embedded in albumen Neck pale to grey. Pallia! roof black, somewhat lighter on gland. Seminal receptacle a broad, folded pouch, about pallial gonoducts. Visceral coil black. 50% of length of bursa copulatrix, usually overlapping Ctenidial filaments 25, medium height, width; ctenid- proximal bursa duct. Seminal receptacle duct medium ium ending slightly anterior to pericardium. Osphradium length. medium-sized, centered slightly posterior to middle of cten- Testis 2.0 whorls, overlapping stomach to posterior edge

Figure 2 Scanning electron micrographs of opercula and radula of P. diablensis Hershler, sp. nov., USNM 883791. A-C. Opercula; bars = 0.43 mm. D. Central radular teeth, bar =15 /urn. E. Lateral teeth, bar = 20 Mm. F. Inner marginal teeth, bar = 30 pm. G. Outer marginal teeth, bar = 20 pm. Page 348 The Veliger, Vol. 38, No. 4 R. Hershler, 1995 Page 349

of style sac. Prostate gland fat bean-shape; oval in cross Etymology: From classical Greek eremos, meaning solitary section; pallial section short. Pallial vas deferens with strong or lonely, and referring to occurrence of this species in the proximal twist. Penis (Figure 3B) large; base broadly rect- fairly remote Smoke Creek Desert and adjacent environs. angular, edges with several weak folds proximally; fila- Diagnosis: Shell broad- to narrow-conic, small- to medi- ment about two-thirds length of base, medium width, gent- um-sized, umbilicate. Penis a narrow blade; penial fila- ly tapering, parallel or slightly oblique to long axis of base; ment medium-elongate; lobe absent. Penial ornament ab- lobe hemispherical, short. Terminal gland medium size, sent. circular or elongate-transverse, usually restricted to ventral surface, rarely interrupted (two units). Penial gland short, Description: Shell (Figures 5B, 7) broad- to narrow-conic; medium width, restricted to base of filament. Filament height 1.7-3.2 mm; whorls 3.75-4.75. Protoconch (Figure darkly pigmented internally; pigment continuing weakly 7K) about 1.5 whorls, early portion densely wrinkled and onto distal penis. lined with scattered spiral striae. Teleoconch whorls convex, strongly shouldered; sculpture of weak growth lines Type locality: Unnamed creek, Del Puerto Canyon, Del and faint spiral striae. Aperture ovate, broadly adnate to Puerto Road, 20 km west of HW 5, Stanislaus County, slightly separated from body whorl. Inner lip complete, California, T. 6 S, R. 6 E, NE VA sec. 8 (Figure 4A). thin to slightly thickened, without columellar shelf. Outer Holotype, USNM 860645; paratypes, USNM 883791, lip thin, orthocline to slightly prosocline. Umbilicus nar- collected by R. Hershler, 4 May 1994. Snails were com- rowly rimate to deeply perforate. Periostracum tan to monly found in this medium-sized (ca. 2 m wide, 0.5 m brown. deep), poorly shaded stream, which was slightly disturbed Operculum (Figure 8A-C) ovate, amber except for or- from pastoral and recreational activities. ange nuclear region; nucleus slightly eccentric; dorsal sur- Remarks: The presence of a penial gland (Pg) in this face frilled. Attachment scar thickened all around, es- species suggests affinity with a large group of western pecially along inner edge. Callus sometimes well devel- American species defined, in part, by this synapomorphy, oped. and closely conforming to Taylor's (1987) " californiensis Buccal mass medium-sized; radular sac protruding be- series" (see Hershler, 1994). Of the members of this group, hind buccal mass as short coil. Radula ribbon elongate, however, only this species plus P. longae and P. taylori about 0.59 x 0.09 mm, with about 65 rows of teeth. (both described below) lack a ventral gland (or any vestige Central radular tooth (Figure 8A, B) trapezoidal, with thereof). Pyrgulopsis diablensis is similar to P. longae in medium to highly indented dorsal edge; lateral cusps 5-6; configuration of the distal female genitalia (i.e., broad coiled central cusp spoonlike, slightly broader and considerably oviduct, frequently pyriform or triangular bursa copu- longer than laterals; basal cusps 1, medium-sized, with latrix), but differs in its more attenuate shell, stronger medium dorsal support. Basal process narrow; basal sock- opercular attachment scar, stouter penis, and larger ter- ets deep. Lateral margins slightly thickened, with pro- minal gland. nounced neck. Lateral tooth (Figure 8F-H) formula 3(4)- This species is thus far known only from the type locality 1-5; neck weakly developed; basal tongue weakly devel- in the Diablo Range, northern San Joaquin Valley (Figure oped; outer wing about 200% of cutting edge length. Mar- 6). ginal teeth (Figure 8F, G, I) with about 21-23 (inner) and 23-28 (outer) cusps; cusps extending onto outer edge of teeth. Stomach caecum small. Pyrgulopsis eremica Hershler, sp. nov. Cephalic tentacles medium to black, sometimes pale distal ly. Snout medium-(more commonly) black. Foot pale to (Figures 5B, 7-9) dark along sides, anterior and posterior edges usually dark. Flurninicola modoci [in part] Hannibal, 1912:187 (Fritter's Opercular lobe broadly darkened along sides, somewhat spring, head of Willow Creek, Honey Lake basin; Trox- lighter along inner edge. Neck pale to dark. Pallial roof, el's spring, Eagle Lake). visceral coil black.

Figure 4 Photographs of type localities of new species of Pyrgulopsis from California. A. Unnamed creek, Del Puerto Canyon, Stanislaus County (P. diablensis Hershler, sp. nov.). B. Unnamed springs tributary to Willow Creek, Lassen County (P. eremica Hershler, sp. nov.). C. Unnamed springs west of Fee Reservoir, Modoc County (P. gibba Hershler, sp. nov.). D. Grapevine Creek, Kern County (P. greggi Hershler, sp. nov.). E. Unnamed spring, ca. 4.8 km west-southwest of Hallelujah Junction, Lassen County (P. longae Hershler, sp. nov.). F. Unnamed spring, 7.4 km south of HW 29 along Seigler Canyon Road, Lake County (P. ventricosa Hershler, sp. nov.). G. Unnamed spring, 4.8 km north of San Luis Obispo, east of HW 101, San Luis Obispo County (P. taylori Hershler, sp. nov.). Page 350 The Veliger, Vol. 38, No. 4

Figure 5 Shells (holotypes) of new Pyrgulopsis species. A. P. diablensU Hershler, sp, nov., USNM 860645 (3,1 mm tall). B. P. eremica Hershler, sp. nov,, USNM 860644 (1.9 mm). C. P. gibba Hershler, sp. nov., USNM 860643 (3.1 mm), D. P. greggi Hershler, sp. nov., USNM 860641 (2.4 mm). E. P. longae Hershler, sp. nov., USNM 860642 (3.0 mm). F. P. taylori Hershler, sp. nov., USNM 860646 (2.0 mm). G. P. ventrkosa Hershler, sp. nov., USNM 860647 (2.4 mm).

Ctenidial filaments 14, short, narrow; ctenidium ending broadly overlapping coils. Distal coil sometimes lightly slightly annTuir to pericardium. Osphnidium medium- pigmented. Oviduct and bursal duct join just behind pallial sized, centered slightly posterior to middle of ctenidial axis. wall. Bursa copulatrix clublike, slightly less than half of Kidney with medium pallial bulge; renal gland longitu- length of albumen gland, narrow relative to width of al- dinal; kidney opening opaque. Rectum straight, broadly bumen gland, with one-fifth to one-half of length posterior overlapping genital ducts. Hypobranchial gland thin; over- to gland. Bursa duct narrow, about as long as bursa cop- lapping rectum, pallial gonoducts. ulatrix, slightly embedded in albumen gland. Seminal re- Distal female genitalia shown in Figure 9A. Ovary 0.75 ceptacle a narrow pouch, slightly less than one-half length whorl, overlapping posterior stomach chamber. Pallial al- of bursa copulatrix, overlapping anterior one-half of bursa bumen gland medium-sized. Capsule gland slightly shorter copulatrix. Seminal receptacle duct medium length. than albumen gland. Genital aperture a broad slit with Testis 2,0 whorls, overlapping stomach to posterior edge short vestibule. Coiled oviduct of two oblique-circular, of style sac. Prostate gland bean-shaped; narrow in cross R. Hershler, 1995 Page 351 section; pallial section medium. Pallial vas deferens with proximal twist. Penis (Figure 9B) small, narrow; base rectangular, smooth along inner edge; filament slightly shorter to slightly longer than base, slightly narrower than base, gently tapering, parallel to long axis of base; lobe absent. Penial ornament absent. Virtual entirety of filament and distal base darkly pigmented internally. Type locality: Unnamed springs tributary to Willow Creek, Willow Creek Valley, Lassen County, California, T. 32 N, R. 11 E, SE % sec. 35 (Figure 4B). Holotype, USNM 860644 (Figure 5B); paratypes, USNM 858264, collected by R. Hershler and D. Sada, 4 August 1990. Snails were commonly found in watercress in this series of small, cool, relatively pristine springs draining to a small meadow alongside Willow Creek. Remarks: This species and P. greggi, described below, differ from all other described members of the genus in lacking penial glands. I interpret this character state as secondarily reduced from a glandular condition, and given that these taxa conform to "typical" Pyrgulopsis in all other respects, place them in this genus rather than create a new higher taxon. It is likely that these species are allied to the group of western American forms having a small distal lobe and weakly developed terminal gland (described in Hershler, 1994): both the lobe and gland are extremely Figure 6 reduced and even sometimes absent in some members of this group (i.e., P. bryantuialkeri Hershler, P. stearnsiana Map showing distributions of P. diablensis Hershler, sp. nov., [Pilsbry], and P. thompsoni Hershler). Pyrgulopsis eremica P. eremica Hershler, sp. nov., P. longae Hershler, sp. nov., and P. taylori Hershler, sp. nov. Symbols may refer to more than differs from P. greggi in its larger size, stronger basal one locality. tongue on lateral radular teeth, smaller ctenidium, larger albumen gland, and occasional pigmentation of coiled oviduct. 6). This snail probably also occurs in additional drainages Both this species and P. longae (described below), which within the above areas (Snowstorm Creek, northeast flank also occurs in the Honey Lake basin, differ from P. melina of Skedaddle Mountains, drainage west-northwest of Ob- Taylor, 1981 (in Taylor & Smith, 1981), described from servation Peak, Van Loan Creek) that could not be thor- Pliocene Honey Lake fossils, by their much smaller size, oughly searched because of access problems. more convex whorls, and weaker columellar callus. Pyrgulopsis eremica is distributed among springs and Material examined: CALIFORNIA. LASSEN COUN- spring brooks within the Great Basin of northeast Cali- TY: unnamed springs, Murrers Lower Meadow, Willow fornia, including a portion of the Lahontan system (Honey Creek Valley, T. 32 N, R. 11 E, SW % sec. 35, USNM Lake basin, Smoke Creek Desert) and smaller basins to 874026; unnamed spring east of Troxel Point, Eagle Lake the north (Eagle Lake basin, Horse Lake basin; Figure basin, T. 32 N, R. 12 E, SW % sec. 5, USNM 858265;

Figure 7 Scanning electron micrographs of shells of P. eremica Hershler, sp. nov. A-C. USNM 858264. D. USNM 858270. E, F. USNM 858267. G, H. USNM 858265. I, J. USNM 858271. K. Protoconch, USNM 858264, bar = 100 /xm. Shell "A" is 2.7 mm tall; other shells printed to same scale.

Figure 8 Scanning electron micrographs of opercula and radula of P. eremica Hershler, sp. nov., USNM 858264. A-C. Opercula, bars = 250 /im, 231 pm, respectively ("B" same scale as "A"). D, E. Central radular teeth, bars = 10 jum, 13.6 fim, respectively. F. Lateral, inner marginal teeth, bar = 23.1 yum. G. Section of radular ribbon, bar = 38 Mm. H. Lateral teeth, bar = 17.6 (im. I. Outer marginal teeth, bar = 20 /xm. Page 352 The Veliger, Vol. 38, No. 4 R. Hershler, 1995 Page 353 Page 354 The Veliger, Vol. 38, No. 4

873210, USNM 874917; unnamed spring, Karlo Road, Secret Valley, T. 31 N, R. 15 E, NE % sec. 3, USNM 874912; Sage Hen Spring, Smoke Creek Desert, T. 33 N, R. 16 E, NE >/4 sec. 35, USNM 858270; unnamed spring east of Sage Hen Spring, T. 33 N, R. 16 E, SW V4 sec. 25, USNM 873411; unnamed spring southwest of Sage Hen Spring, T. 33 N, R. 16 E, NE % sec. 35, USNM 858269; Big Spring, Smoke Creek Desert, T. 33 N, R. 16 E, NW Vi sec. 1, USNM 858271, USNM 873392; unnamed springs, Shinn Ranch, Smoke Creek Desert, T. 33 N, R. 16 E, SW 1/4 sec. 36, USNM 858268, USNM 873403; unnamed spring, east of Rush Creek Ranch, Smoke Creek Desert, T. 31 N, R. 17 E, SE !4 sec. 11, USNM 858266.

Pyrgulopsis gibba Hershler, sp. nov. (Figures 5C, 10-12) Etymology: From Latin gibber, meaning swollen, and referring to enlarged penial lobe in this species. B Diagnosis: Shell ovate- to narrow-conic, medium-sized, umbilicate. Penial filament short; lobe enlarged. Penial ornament a terminal gland (often interrupted), Dg3 (sometimes absent), and ventral gland. Description: Shell (Figures 5C, 10) ovate- to narrow- conic; height 2.7-3.5 mm; whorls 4.0-4.75. Protoconch (Figure 10H) about 1.5 whorls, early portion weakly wrinkled and incised with weak spiral grooves. Teleoconch whorls convex, slightly shouldered; sculpture of medium growth lines. Aperture ovate, narrowly adnate to or slightly separated from body whorl. Inner lip complete, thin, without columellar shelf. Outer lip thin, orthocline to slightly prosocline. Umbilicus narrowly rimate (near ab- Figure 9 sent) to perforate. Periostracum tan-brown. Genital morphology of P. eremica Hershler, sp. nov., USNM Operculum (Figure 11A-C) ovate, amber except for 858264. A. Distal female genitalia (left side). B. Penes (dorsal orange streaks in nuclear region; nucleus slightly eccentric; aspect). Bar = 0.5 mm. Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov dorsal surface weakly frilled. Attachment scar margin often = coiled oviduct, Sr = seminal receptacle, Vc = ventral channel. broadly thickened all around. Callus well developed. Buccal mass medium-sized; radular sac protruding behind buccal mass as short coil. Radula ribbon elongate, unnamed springs northeast of Horse Lake, Horse Lake about 0.86 x 0.15 mm, with about 65 rows of teeth. basin, T. 33 N, R. 13 E, NE % sec. 14, USNM 873138; Central radular tooth (Figure 1 ID) trapezoidal, with highly Tule Patch Spring, Secret Valley, T. 32 N, R. 15 E, SE indented dorsal edge; lateral cusps 5; central cusp spoon- !/4 sec. 10, USNM 854569, USNM 858267, USNM like, slightly broader and considerably longer than laterals; 874053; unnamed spring, northern Secret Valley, T. 31 basal cusps 1, short, with weak dorsal support. Basal pro- N, R. 15 E, NE i/4 sec. 3, USNM 858272; Sellicks Springs, cess medium width; basal sockets deep. Lateral margins Secret Valley, T. 31 N, R. 15 E, NW % sec. 7, USNM thickened, with medium to pronounced neck. Lateral tooth

Figure 10 Scanning electron micrographs of shells of P. gibba Hershler, sp. nov. A-E. USNM 858275. F. USNM 858274. G. USNM 858273. H. Protoconch, USNM 858275, bar = 136 Mm. Shell "A" is 2.7 mm; other shells printed to same scale. R. Hershler, 1995 Page 355 Page 356 The Veliger, Vol. 38, No. 4 R. Hershler, 1995 Page 357

Cgo

Figure 12 Genital morphology of P. gibba Hershler, sp. nov. A. Distal female genitalia (left side), USNM 858275, bar = 0.5 mm. B. Penis (dorsal aspect, left; ventral aspect, right), USNM 858275, bar = 0.5 mm. C. Ibid., USNM 858273. D. Ibid., USNM 858274. Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled oviduct, Dg3 = dorsal gland, Sr = seminal receptacle, Tg = terminal gland, Vc = ventral channel, Vg = ventral gland.

(Figure HE, F) formula 3-1-1; neck weakly developed; Snout medium to dark; distal lips sometimes pale. Foot basal tongue well developed; outer wing about 170% of pale to light along sides; anterior/posterior edges some- cutting edge length. Marginal teeth (Figure 11E-G) with times medium to dark. Opercular lobe dark along perim- about 23-36 (inner) and 28 (outer) cusps; cusps extending eter. Neck near pale to dark. Pallial roof, visceral coil onto outer edge of teeth. Stomach caecum small. black. Cephalic tentacles near pale to dark brown to black. Ctenidial filaments 18-20, broad and tall; ctenidium

Figure 11 Scanning electron micrographs of opercula and radula of P. gibba Hershler, sp. nov., USNM 858275. A-C. Opercula, bars = 0.33 mm, 0.38 mm, respectively ("C" same scale as "B"). D. Central radular teeth, bar = 17.6 (im. E. Lateral, inner marginal teeth, bar = 25 pm. F. Lateral, marginal teeth, bar = 38 (im. G. Outer marginal teeth, bar = 27 ^m. Page 358 The Veliger, Vol. 38, No. 4 slightly overlapping pericardium posteriorly. Osphradium Dgl (although a very weak papule, perhaps conforming small, centered slightly posterior to middle of ctenidial axis. to this gland, rarely is present). Kidney with medium pallial bulge; renal gland longitu- This snail occurs in the Great Basin of northeastern dinal; kidney opening white. Rectum straight, broadly California (northern Surprise Valley, Duck Lake Valley; overlapping genital ducts. Hypobranchial gland thin; over- Figure 13). lapping rectum, pallial gonoducts, narrow portion of pos- Material examined: CALIFORNIA. LASSEN COUN- terior pallial roof. TY: Unnamed spring, Old Marr Ranch, Tuledad Canyon, Distal female genitalia shown in Figure 12A. Ovary Duck Flat, T. 37 N, R. 17 E, NE % sec. 31, USNM 0.5-0.75 whorl, overlapping posterior stomach chamber. 858273. MODOC COUNTY: Unnamed spring, north- Pallial albumen gland short. Capsule gland slightly shorter west side of Lake Annie, northwest of Lake Annie Road, than albumen gland. Genital aperture a broad slit on slightly Surprise Valley, T. 47 N, R. 16 E, SW VS sec. 26, USNM raised papilla, with short vestibule. Coiled oviduct lightly 858274. pigmented, of two short, oblique-circular, slightly overlapping coils. Oviduct and bursal duct join just behind Pyrgulopsis greggi Hershler, sp. nov. pallial wall. Bursa copulatrix broadly ovate or saclike, slightly less than one-half of length of albumen gland, and (Figures 5D, 14-16) slightly more than half of width of gland, with about 10- Amnicola.—Cooper, 1869:217 (In and about Ft. Tejon). 30% of length posterior to gland. Bursa duct narrow, about one-half of length of bursa copulatrix, well embedded in Etymology: This species is named in honor of the late albumen gland. Seminal receptacle sometimes a slender Wendell O. Gregg, in recognition of his extensive research pouch, sometimes lightly pigmented, about 20% of length on the genus Pyrgulopsis in California and for his recog- of bursa copulatrix, overlapping or lateral to anterior por- nizing the distinctiveness of this snail when he collected it tion of bursa copulatrix. Seminal receptacle duct medium in 1964. length. Diagnosis: Shell conical, small, umbilicate. Penis simple, Testis 1.5 whorls, overlapping posterior stomach cham- bladelike; filament medium length, lobe absent. Penial or- ber. Prostate gland bean-shaped; near-circular in cross nament absent. section; pallial section short. Pallial vas deferens with proximal twist. Penis (Figure 12B-D) large; base folded along Description: Shell (Figures 5D, 14) conical; height, 1.6- inner edge; filament considerably shorter than base, nar- 2.6 mm; whorls 3.75-5.0. Protoconch (Figure 14F) about row, tapering, parallel or slightly angled relative to long 1.5 whorls, early portion punctate and with several widely axis of base; lobe broadly rectangular, about as long as separated spiral lines. Teleoconch whorls convex, shoul- base, often bifurcate or swollen distally. Terminal gland dered; sculpture of medium growth lines. Aperture ovate, elongate-transverse, usually curving onto both dorsal and narrowly adnate to (more commonly) slightly separated ventral surfaces (most of length on latter), often inter- from body whorl. Inner lip complete, thin, without colu- rupted, with units restricted to inner and outer portions mellar shelf. Outer lip thin, orthocline. Umbilicus small, of lobe, sometimes with a third, intermediate (ventral) unit chinklike to broadly rimate. or sometimes fused as single, elongate unit. Dg3 often Operculum (Figure 15A-C) broadly ovate, amber ex- present, either as small papule (sometimes double) or large, cept for orange nuclear region; nucleus slightly eccentric; raised unit. Dorsal surface rarely with a very small prox- dorsal surface frilled. Attachment scar margin often broad- imal gland possibly representing reduced Dgl. Ventral ly thickened all around. Callus well developed. gland large, elongate, borne on pronounced swelling, often Buccal mass medium-sized; radular sac protruding be- accompanied by one to two smaller distal glands. Proximal hind buccal mass as short coil. Radula ribbon elongate, two-thirds of filament darkly pigmented internally. about 0.5 x 0.08 mm, with about 52 rows of teeth. Central radular tooth (Figure 15D) trapezoidal, with highly in- Type locality: Unnamed springs west of Fee Reservoir, dented dorsal edge; lateral cusps 5-7; central cusp pointed, Surprise Valley, Modoc County, California, T. 46 N, R. considerably broader and slightly longer than laterals; bas- 17 E, NE VA sec. 20 (Figure 4C). Holotype, USNM 860643 al cusps 1, medium length, with weak dorsal support. Basal (Figure 5C); paratypes, USNM 858275, collected by R. process narrow; basal sockets deep. Lateral margins thick- Hershler and D. Sada, 9 August 1990. Snails were com- ened, with pronounced neck. Lateral tooth (Figure 15E, monly found in mud and scattered watercress in a series F) formula 4-1-4(5); neck weakly developed; basal tongue of degraded springs draining to a small meadow. well developed; outer wing about 180% of cutting edge Remarks: This species is unique among members of the length. Marginal teeth (Figure 15E-G) each with about genus in having penial ornament of terminal gland, Dg3, 22-26 cusps; cusps extending onto outer edge of teeth. and ventral gland. Affinities of the snail may lie with the Stomach caecum small. group of western American species conforming in part to Cephalic tentacles pale to medium grey to black (on Natricola Gregg & Taylor, 1965, but it differs in lacking proximal half). Snout medium to black. Foot pale, medium R. Hershler, 1995 Page 359

anterior to or slightly overlapping bursa copulatrix. Sem- inal receptacle duct medium length. Testis 2.0 whorls, overlapping entire stomach and ex- • gibba tending to near edge of prostate gland. Prostate gland bean- * greggi shaped, narrow in cross section; pallial section medium. Pallial vas deferens with weak proximal bend. Penis (Fig- O ventricosa ure 16B, C) small; base broadly to narrowly rectangular; filament about two-thirds of length of base, distally tapering, oriented parallel to long axis of base; lobe absent. Filament considerably narrower to sub-equal to base. In- ner curvature of base lined with shallow folds. Proximal two-thirds of filament darkly pigmented internally. Type locality: Grapevine Creek, Fort Tejon State His- torical Park, Castac Valley, Kern County, California, T. 9 N, R. 19 W, NE !4 section 16 (Figure 4D). Holotype, USNM 860641 (Figure 5D); paratypes, USNM 874139, collected by R. Hershler, 10 April 1991. Snails were commonly found in mud and watercress in this medium-sized, moderately impacted stream (ca. 1-2 m wide, 0.25 m deep). Remarks: This species is restricted to Grapevine Creek drainage in the Upper Kern River basin (Figure 13). Material examined: Topotypes, USNM 873418, WBM 4629, WBM 4689; Unnamed spring about 0.8 km north Figure 13 of above, T. 9 N, R. 19 W, NE % sec. 9, USNM 873402, USNM 874140. Map showing distributions of P. gibba Hershler, sp. nov., P. greggi Hershler, sp. nov., and P. ventricosa Hershler, sp. nov. Symbols may refer to more than one locality. Pyrgulopsis longae Hershler, sp. nov. (Figures 5E, 17-19) Etymology: The species name refers to distribution of the grey or black along sides, with pigment especially heavy snail within Long Valley. on anterior and posterior edges. Opercular lobe dark along Diagnosis: Shell ovate- to narrow-conic, medium-sized, margins. Neck pale to medium. Pallial roof, visceral coil umbilicate. Penial filament short, lobe medium-sized. Pen- mottled to uniformly black dorsally; pigment lighter on ial ornament a short penial gland and small terminal gland. genital ducts. Ctenidial filaments 18, medium height and width; cte- Description: Shell (Figures 5E, 17) ovate- to.narrow- nidium slightly overlapping pericardium posteriorly. Os- conic; height 2.2-3.0 mm; whorls 4.0-4.75. Protoconch phradium medium-sized, centered slightly posterior to (Figure 17F) 1.5 whorls, near smooth. Teleoconch whorls middle of ctenidial axis. Kidney with medium pallial bulge; convex, slightly shouldered; sculpture of weak growth lines. renal gland longitudinal; kidney opening white. Rectum Aperture ovate, usually slightly separated from body whorl. straight, broadly overlapping genital ducts. Hypobranchial Inner lip complete, thin, without columellar shelf. Outer gland thin; overlapping rectum, pallial gonoducts. lip thin, orthocline. Umbilicus small, chinklike to broadly Distal female genitalia shown in Figure 16A. Ovary 1.0 rimate. Periostracum dark tan. whorl, partly overlapping posterior stomach chamber. Pal- Operculum (Figure 18A-C) narrowly ovate, amber ex- lial albumen gland very short. Capsule gland slightly shorter cept for reddish nuclear region; nucleus highly eccentric; than albumen gland. Genital aperture a broad slit with dorsal surface weakly frilled. Attachment scar margin often short anterior vestibule. Coiled oviduct of two small, over- well thickened all around. Callus well developed. lapping, circular-oblique loops. Oviduct and bursal duct Buccal mass medium-sized; radular sac protruding be- join just behind pallial wall. Bursa copulatrix ovate to hind buccal mass as short coil. Radular ribbon elongate, clublike, medium length (33%) and width, with 20-25% about 0.65 x 0.10 mm, with about 70 rows of teeth. of length posterior to albumen gland. Bursal duct medium Central radular tooth (Figure 18D, E) trapezoidal, with width, about 150% of length of bursa copulatrix, partly highly indented dorsal edge; lateral cusps 5; central cusp embedded in albumen gland. Seminal receptacle pouchlike, rounded, considerably broader and slightly longer than sometimes folded, about 40% of length of bursa copulatrix, laterals; basal cusps 1, elongate, with moderate dorsal sup- Page 360 The Veliger, Vol. 38, No. 4

Figure 14 Scanning electron micrographs of shells of P. greggi Hershler, sp. nov. A-C. USNM 874139. D, E. USNM 874140. F. Protoconch, USNM 874139, bar = 120 )im. Shell "A" is 2,3 mm tall; other shells printed to same scale.

Figure 15 Scanning electron micrographs of operatla and radula of P. greggi Hershler, sp. nov., USNM 873418. A-C. Opercula, bars = 250 nm, 270 pm, respectively ("C" same scale as "B"). D. Central radular teeth, bar = 10 Mm. E, F. Lateral, inner marginal teeth, bars = 17.6 ^n. G. Outer marginal teeth, bar — 17.6 nm. R. Hershler, 1995 Page 361 Page 362 The Veliger, Vol. 38, No. 4

medium-sized, centered slightly posterior to middle of ctenidial axis. Kidney with medium pallial bulge; renal gland longitudinal; kidney opening white. Rectum straight, broadly overlapping genital duct. Hypobranchial gland medium thickness; overlapping rectum, pallial gonoducts, roof of pallial cavity alongside rectum. Distal female genitalia shown in Figure 19A. Ovary 0.75-1.0 whorl, overlapping posterior stomach chamber. Pallial albumen gland short. Capsule gland shorter than albumen gland. Genital aperture a broad slit without vestibule. Coiled oviduct a small proximal kink or twist fol- lowed by elongate horizontal coil. Oviduct and bursal duct join just behind pallial wall. Bursa copulatrix ovoid-pyriform, medium length and width, with about one-half of length posterior to albumen gland. Bursal duct narrow (broadening distally), about two-thirds of length of bursa copulatrix, partly embedded in albumen gland. Seminal receptacle globular-pyriform, sometimes folded, about one- half of length of bursa copulatrix, overlapping anterior half of bursa copulatrix. Seminal receptacle duct short. Testis 1.25 whorls, overlapping posterior stomach chamber. Prostate gland bean-shaped, narrow in cross section; pallial section very short to absent. Pallial vas deferens without proximal twist. Penis (Figure 19B) large; base broadly rectangular; filament short, tapering distally; lobe medium-sized, tapering distally. Penial gland a short strip on base of filament. Terminal gland small, transverse- circular, usually borne on ventral surface. Proximal half of penial filament with dark internal pigment; distal half of penis base with scattered black granules. Type locality: Unnamed spring about 4.8 km west-southwest of Hallelujah Junction, Long Valley, Lassen County, California, T. 22 N, R. 17 E, SW % sec. 9 (Figure 4E). Figure 16 Holotype, USNM 860642 (Figure 5E); paratypes, USNM Genital morphology of P. greggi Hershler, sp. nov. A. Distal 858262, collected by R. Hershler and D. Sada, 3 August female genitalia (left side), USNM 874139. B, C. Penes (dorsal 1990. Snails were commonly found on stones and in wa- aspect), USNM 874139 (left), USNM 874140 (right). Bar = tercress of this large spring, which was situated adjacent 0.5 mm. Ag = albumen gland, nd, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled oviduct, to a residence and had been recently excavated. Sr = seminal receptacle, Vc = ventral channel. Remarks: This species is known only from the type locality in the southern end of the Honey Lake basin (Figure 6). port. Basal process narrow; basal sockets deep. Lateral margins thickened, with pronounced neck. Lateral tooth (Figure 18E) formula 4-1-4; neck well developed; basal Pyrgulopsis taylori Hershler, sp. nov. tongue well developed; outer wing about 150% of cutting (Figures 5F, 20-22) edge length. Marginal teeth (Figure 18E-G) with about 24 (inner) and 28 (outer) cusps; cusps extending onto outer Etymology: This species is named in honor of Dwight W. edge of teeth. Stomach caecum small. Taylor, in recognition of his many years of fieldwork and Cephalic tentacles with medium grey to black pigment associated research on hydrobiids both in California and proximally. Snout light to medium grey. Foot light along throughout the western United States. sides; anterior edge light to medium. Opercular lobe usually black along margins. Neck pale to light grey. Pallial Diagnosis: Shell ovate- to narrow-conic or pupiform, small, roof, visceral coil uniformly black. umbilicate. Penial filament elongate; lobe short. Penial Ctenidial filaments 22, medium height and width; cte- ornament a terminal gland and occasionally a small penial nidium slightly overlapping pericardium. Osphradium gland. R. Hershler, 1995 Page 363

Figure 17 Scanning electron micrographs of shells of P. hngae Hershler, sp. nov,, USNM 858262. F. Protoconch, bar = 136 /an. Shell "A" is 2.5 mm tall; other shells printed to same scale.

Description: Shell (Figures 5F, 20) ovate- to narrow-conic: cline. Umbilicus small, chinklike or perforate. Periostra- or pu pi form; height 2.0-3.3 mm; whorls 4,0-5.25. Pro- cum tan. toconch (Figure 201) about 1.5 whorls, early portion wrin- Operculum (Figure 21A-C) narrowly ovate, very thin, kled, with a few weak spiral striae. Teleoconch whorls light amber; nucleus highly eccentric; dorsal surface weak- near flat to medium convexity, narrowly shouldered; sculp- ly frilled. Attachment scar usually only weakly thickened ture of strong growth lines crossed by very weak spiral along inner edge. Callus weakly developed. striae. Aperture ovaie-pyriform, often enlarged, broadly Buccal mass medium-sized; radular sac protruding be- adnate to slightly separated from body whorl. Inner lip hind buccal mass as short coil. Radula ribbon moderately complete, thin or slightly thickened, columellar shelf often elongate, about 0.9 x 0.09 mm, with about 60 rows of pronounced. Outer lip thin or slightly thickened, ortho- teeth. Central radular tooth (Figure 21D) trapezoidal, Page 364 The Veliger, Vol. 38, No. 4 R. Hershler, 1995 Page 365

with well-indented dorsal edge; lateral cusps 5; central cusp pointed, considerably broader and longer than laterals; basal cusps 1, short, with weak dorsal support. Basal process narrow; basal sockets deep. Lateral margins thickened, with medium neck. Lateral tooth (Figure 21E) formula 3-1-3(4); neck very weakly developed; outer wing 150-180% of cutting edge length. Marginal teeth (Figure 21F, G) with about 17-22 (inner) and 19-20 (outer) cusps; cusps extending onto outer edge of teeth. Stomach caecum small. Cephalic tentacles pale or with light grey patch proximally. Snout pale to medium grey. Foot pale or with light grey patch along anterior edge. Opercular lobe black along inner edge. Neck pale or very light grey. Pallial roof, visceral coil variably pigmented brown to black, rarely uniformly so. Ctenidial filaments about 17, short, narrow, weakly pleated; ctenidium not overlapping pericardium posteriorly. Osphradium medium-sized, positioned centrally or slightly posterior to middle of ctenidial axis. Kidney with medium-sized pallial bulge; renal gland longitudinal; kidney opening white. Rectum straight, broadly overlapping genital ducts. Hypobranchial gland thin; overlapping rectum, pallial gonoducts. Distal female genitalia shown in Figure 22A. Ovary 0.5-0.75 whorl, slightly overlapping posterior stomach chamber. Pallial albumen gland short. Capsule gland slightly shorter than albumen gland. Genital aperture a large subterminal pore, with very short vestibule. Coiled oviduct a small, near circular coil preceeded by proximal twist. Oviduct and bursal duct join well behind pallial wall. Bursa copulatrix simply ovate, up to one-third length of albumen gland, and one-half width of gland, with about 50-65% of length posterior to gland. Bursa duct medium width, well embedded in albumen gland. Seminal receptacle pouchlike, sometimes folded, 50-70% of length of bursa copulatrix, partly overlapping anterior portion of bursa copulatrix. Seminal receptacle duct medium length. Testis 1.5-2.0 whorls, overlapping posterior and portion of anterior stomach chambers. Prostate gland narrowly bean-shaped; oval in cross section; pallial section medium Figure 19 length. Pallial vas deferens with weak proximal bend. Genital morphology of P. longae Hershler, sp. nov., USNM Penis (Figure 22B, C) medium-sized; edges unfolded; fil- 858262. A. Distal female genitalia (left side). B. Penis (dorsal ament sometimes as long as base, narrow, tapering, parallel aspect, left; ventral aspect, right). Bar = 0.5 mm. Ag = albumen to strongly oblique to long axis of base; lobe hemispherical, gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled oviduct, Pg = penial gland, Sr = short (rarely near absent). Terminal gland circular or seminal receptacle, Tg = terminal gland, Vc = ventral channel. ovate, usually positioned on ventral surface. Penial gland small (often absent), near circular, positioned near base of filament. Filament darkly pigmented internally.

Figure 18 Scanning electron micrographs of opercula and radula of P. longae Hershler, sp. nov., USNM 858262. A-C. Opercula; bars = 0.33 mm, 0.38 mm, 0.30 mm, respectively. D. Central radular teeth, bar =15 fim. E. Central, lateral, marginal teeth, bar = 25 fim. F, G. Marginal teeth, bars = 23.1

# E

H R. Hershler, 1995 Page 367

Type locality: Unnamed spring tributary to San Luis narrow. Periostracum tan; shell usually covered with thick Obispo Creek, 4.8 km north of San Luis Obispo, east of organic deposits. HW 101, San Luis Obispo County, California, T. 30 S, Operculum (Figure 24A-C) ovate, amber with reddish R. 13 E, SE VA sec. 7 (Figure 4G). Holotype, USNM center of attachment scar; nucleus eccentric; dorsal surface 860646 (Figure 5F); paratypes, USNM 883792, collected frilled. Attachment scar margin broadly thickened all by R. Hershler, 6 May 1994. Snails were found in mod- around. Callus well developed. erate abundance on rocks and in leaf litter along this small Buccal mass medium-sized; radular sac protruding be- spring brook, which was in fairly good condition despite hind buccal mass as short coil. Radula ribbon moderately some recreational impacts and historic diversion near the elongate, about 0.65 x 0.11 mm, with about 50 rows of source. teeth. Central radular tooth (Figure 24D) trapezoidal, with medium to highly indented dorsal edge; lateral cusps Remarks: This snail is known only from San Luis Obispo 4-5; central cusp pointed, considerably broader and longer Creek drainage along the south-central California coast than laterals; basal cusps 1, short, with weak dorsal sup- (Figure 6). port. Basal process narrow; basal sockets deep. Lateral Material examined. CALIFORNIA. SAN LUIS OBIS- margins weakly thickened, with weak neck. Lateral tooth PO COUNTY: topotypes, USNM 874459, WBM 3865; (Figure 24E) formula 3-1-4; neck weakly developed; basal Unnamed spring tributary to Brizziolari Creek, 1.6 km tongue well developed; outer wing 160-180% of cutting north of California Polytechnic State University, San Luis edge length. Marginal teeth (Figure 24E-G) with about Obispo, T. 30 S, R. 12 E, unsurveyed, USNM 883789; 16-20 (inner) and 20-23 cusps; cusps extending onto outer Chorro Creek, Camp San Luis Obispo, T. 30 S, R. 12 E, edge of teeth. Stomach caecum small. NW Y* sec. 3, USNM 854590; Unnamed spring tributary Cephalic tentacles light grey to black; pigment some- to Morro Creek, west side of HW 41, T. 29 S, R. 11 E, times a longitudinal band on pale background. Snout grey SE % sec. 2, USNM 883788. to black, rarely red. Foot grey to black along anterior end, sometimes similarly pigmented along rest of margin, oth- erwise pale. Opercular lobe black along inner edge, lighter along rest of margin. Neck pale to near black. Pallial roof mottled to near uniform black; visceral coil slightly lighter. Pyrgulopsis ventricosa Hershler, sp. nov. Ctenidial filaments 22, medium width, tall; ctenidium slightly overlapping pericardium posteriorly. Osphradium (Figures 5G, 23-25) small, centered well posterior to middle of ctenidial axis. Pomatiopsis intermedia.—Cooper, 1876:36 ("near Clear Kidney with medium pallial bulge; renal gland longitu- Lake"). dinal; kidney opening white. Rectum straight, broadly Etymology: From Latin ventricosus, meaning bulging, and overlapping genital ducts. Hypobranchial gland thin; over- referring to the well-developed ventral penial glands in lapping rectum, pallial gonoducts. this species. Distal female genitalia shown in Figure 25A. Ovary 1.0 whorl, abutting posterior edge of stomach. Pallial albumen Diagnosis: Shell ovate- to narrow-conic, medium-sized, gland short. Capsule gland about as long as or slightly umbilicate. Penial filament, lobe medium-sized. Penial or- longer than albumen gland. Genital aperture a broad slit; nament a transverse terminal gland, elongate penial gland, vestibule absent or very weakly developed. Coiled oviduct elongate Dgl, short Dg2, variable Dg3, an additional elon- circular-horizontal, simple or kinked near mid-point. Ovi- gate dorsal gland, and two prominent ventral glands. duct and bursal duct join slightly behind pallial wall. Bursa copulatrix ovate, sometimes with blunt anterior edge, 80- Description: Shell (Figures 5G, 23) ovate- to narrow- 110% length of albumen gland, as wide or slightly nar- conic; height 2.2-2.6 mm; whorls 4.0-4.5. Protoconch rower than albumen gland, with almost entire length pos- (Figure 23F) about 1.25 whorls, early portion strongly terior to gland. Bursal duct narrow, short (about one- wrinkled with occasional spiral lines. Teleoconch whorls fourth to one-third length of bursa copulatrix), weakly convex, strongly shouldered; sculpture of weak growth embedded in albumen gland. Seminal receptacle a small lines and faint spiral striae. Aperture ovate, adnate to well pouch, less than one-fourth length of bursa copulatrix, separated from body whorl. Inner lip complete, thin, with- overlapping anterior portion of bursa copulatrix. Seminal out columellar shelf. Outer lip thin, orthocline. Umbilicus receptacle duct medium length.

Figure 20 Scanning electron micrographs of shells of P. taylori Hershler, sp. nov. A-E, USNM 883792. F-H, USNM 883789. I. Protoconch, USNM 883792, bar =136 nm. Shell "A" is 2.2 mm tall; other shells printed to same scale. Page 368 The Veliger, Vol. 38, No. 4 R. Hershler, 1995 Page 369

Figure 22 Genital morphology of P. taylori Hershler, sp. nov., USNM 883792. A. Distal female genitalia (left side). Bar = 0.5 mm. B. C. Penes (dorsal aspect, left; ventral aspect, right). Bar = 0.5 mm. Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled oviduct, Pg = penial gland, Sr = seminal receptacle, Tg = terminal gland, Vc = ventral channel.

Testis, 1.5 whorls, overlapping posterior and portion of gland transverse, usually interrupted into two distinct units anterior stomach chambers. Prostate gland bean shape; (rarely with third intermediate glandular dot), usually narrow oval in cross section; pallial section short. Pallial coursing over dorsal and ventral surfaces. Penial gland vas deferens with very large proximal loop. Penis (Figure elongate, covering most of length of filament. Dgl elongate, 25B-D) large; base rectangular, weakly folded along inner extending along length of base near outer edge, sometimes edge; filament considerably shorter than base, medium fused with either Dg3 or additional dorsal glands. Dg2 width, tapering, slightly oblique to long axis of base; lobe short. Dg3 dotlike to large, borne on pronounced swelling. slightly tapering distally, shorter than filament. Terminal Dorsal penis also with an additional elongate gland (as

Figure 21 Scanning electron micrographs of opercula and radula of P. taylori Hershler, sp. nov., USNM 883792. A-C. Opercula; bars = 231 /im, 250 pm. "C" printed to same scale as "A." D. Central radular teeth, bar = 12 jim. E. Lateral teeth, bar = 15 (im. F. Inner marginal teeth, bar = 12 fim. G. Outer marginal teeth, bar = 17.6 fim. Page 370 The Veliger, Vol. 38, No. 4

Figure 23 Scanning electron micrographs of shells of P. ventricosa Hershler, sp, nov., USNM 883790. F, Protoconch, bar 120 jim. Shell "A" is 2.9 mm tall; other shells printed to same scale.

Figure 24 Scanning electron micrographs of opercula and radula of P. ventricosa Hershler, sp, nov., USNM 883790. A-C. Opercula, bar = 0,30 mm. D. Central radular teeth, bar = 12 jim. E. Central, lateral, inner marginal teeth, bar — 17.6 jim. F. Inner marginal teeth, bar = 17.6 jim. G. Inner and outer marginal teeth, bar = 23,1 jim. R. Hershler, 1995 Page 371 Page 372 The Veliger, Vol. 38, No. 4

B Dg3

Dg2

Figure 25 Genital morphology of P. ventricosa Hershler, sp. nov., USNM 883790. A. Distal female genitalia (left side). Bar = 0.5 mm. B. Penis (dorsal aspect, left; ventral aspect, right). Bar = 0.5 mm. C, D. Penes (dorsal aspects). Scale as in "B." Ag = albumen gland, Bu = bursa copulatrix, Cg = capsule gland, Cgo = capsule gland opening, Cov = coiled oviduct, Dgl = gland along right edge, Dg2 = gland along left distal edge, Dg3 = gland along right edge of lobe, Pg = penial gland, Sr = seminal receptacle, Tg = terminal gland, Vc = ventral channel, Vg = ventral gland. well as one to two small glandular dots) centrally posi- 5G); paratypes, USNM 883790, collected by R. Hershler tioned between Dgl and Dg3, apparently sometimes fused and S. Ellis, 3 May 1994. Snails were commonly found with former. Ventral penis with distal transverse gland on vegetation in this small (1 m wide, 3 cm deep), spring- (rarely interrupted into two units) borne on low swelling fed stream, which had been diverted and additionally im- and large, near central gland borne on tall swelling. Fil- pacted by residential activities. ament darkly pigmented internally along most of length. Remarks: This species is similar to the most derived mem- Type locality: Unnamed creek, Seigler Canyon, 7.4 km bers of the "californiensis series" (see above), which are south of HW 29 along Seigler Canyon Road, Clear Lake characterized in part by possession of a full complement basin, Lake County, California, T. 12 N, R. 7 W, NE % of glands on the penis (Pg, Tg, Dgl-3) and an enlarged sec. 19 (Figure 4F). Holotype, USNM 860647 (Figure bursa copulatrix. Of the members of this group, only P. R. Hershler, 1995 Page 373

californiensis, from coastal regions along southern Califor- Sada. I thank Barry Roth and an anonymous reviewer for nia, and P. wongi, from Owens River drainage and several comments on the manuscript. other basins to the east of Sierra Nevada, share with our species two well-developed ventral glands (Vg) and a very small seminal receptacle. Pyrgulopsis ventricosa shares with LITERATURE CITED

P. californiensis an ovate-narrow conic shell and short bur- ARNOLD, R. 1909. Paleontology of the Coalinga District, Fres- sal duct, but is distinguished by penial features, including no and Kings counties, California. United States Geological the generally longitudinal (not transverse) orientation of Survey, Bulletin 396:1-173. Dgl, shorter Dg2, fewer and smaller additional dorsal COOPER, J. G. 1869. On the distribution and localities of West glands, and more pronounced proximal ventral gland. Coast Helicoid land shells, &C. American Journal of Con- chology 4:211-240. Late Cenozoic hydrobiids from the Cache Formation in COOPER, J. G. 1876. On shells of the west slope of North the Clear Lake basin have been provisionally allocated to America. No. III. Proceedings of the California Academy of Hydrobia andersoni Arnold, 1909 (Taylor, 1966 and ref- Sciences 6:14-27. erences cited therein; Rymer et al., 1988), which in turn GREGG, W. O. & D. W. TAYLOR. 1965. Fontelicella (Proso- was described from fossils of the Tulare Formation in branchia: Hydrobiidae), a new genus of West American Kettleman Hills, located near the southern end of San freshwater snails. Malacologia 3:103-110. Joaquin Valley. Although I have not seen Cache For- HANNIBAL, H. 1912. A synopsis of the Recent and Tertiary freshwater Mollusca of the California Province, based upon mation material and thus cannot ascertain possible con- an ontogenetic classification. Proceedings of the Malacolog- specificity with our snail, this novelty is clearly distin- ical Society of London 10:112-166, 167-211. guished from typical H. andersoni by its smaller, squatter HERSHLER, R. 1989. Springsnails (Gastropoda: Hydrobiidae) shell and shallower sutures. of Owens and Amargosa River (exclusive of Ash Meadows) This species is restricted to Seigler Creek drainage in drainages, Death Valley system, California-Nevada. Pro- the south end of the Clear Lake basin (Figure 13). Al- ceedings of the Biological Society of Washington 102:176- 248. though unpublished historic records suggested that the HERSHLER, R. 1994. A review of the North American fresh- snail was formerly widespread in this region (at least until water snail genus Pyrgulopsis (Hydrobiidae). Smithsonian the 1970s), I was able to locate only one other living colony Contributions to Zoology 554:1-115. a short distance from the type locality in 1994. HERSHLER, R. & W. L. PRATT. 1990. A new Pyrgulopsis from Southeastern California, with a model for historical devel- Material examined: CALIFORNIA. LAKE COUNTY: opment of the Death Valley hydrographic system. Proceed- Unnamed spring, Seigler Canyon, T. 12 N, R. 7 W, NE ings of the Biological Society of Washington 103:279-299. '/4 sec. 19, USNM 873134; Seigler Springs, T. 12 N, R. HERSHLER, R. & D. W. SADA. 1987. Springsnails (Gastropoda: 8 W, NE % sec. 24, USNM 874925. Hydrobiidae) of Ash Meadows, Amargosa Basin, Califor- nia-Nevada. Proceedings of the Biological Society of Wash- ington 100:776-843. MINCKLEY, W. L., D. A. HENDRICKSON & C. E. BOND. 1986. Geography of western North American freshwater fishes: ACKNOWLEDGMENTS description and relationships to intracontinental tectonism. I thank F. G. Hochberg and P. Scott (Santa Barbara Pp. 519-613 [+ bibliography] in C. H. Hocutt & E. O. Museum of Natural History) for lending specimens under Wiley (eds.), Zoogeography of North American Freshwater Fishes. J. Wiley & Sons: New York. their care. K. Cummings (Illinois Natural History Sur- RYMER, M. J., B. ROTH, J. P. BRADBURY & R. M. FORRESTER. vey), J. Landye, and W. Miller (University of Arizona) 1988. Depositional environments of the Cache, Lower Lake, generously donated specimens and notes. and Kelseyville Formations, Lake County, California. Pp. Scanning electron micrographs were taken by Susann 45-61 in J. D. Sims (ed.), Late Quaternary Climate, Tec- Braden of the NMNH (USNM) Scanning Electron Mi- tonism, and Sedimentation in Clear Lake, Northern Cali- croscopy Laboratory, and prints of these were made by fornia Coast Ranges. Geological Society of America Special Paper 214:1-225. Victor Krantz, and staff of the NMNH Office of Printing TAYLOR, D. W. 1966. Summary of North American blancan and Photographic Services. Shell drawings and drainage nonmarine mollusks. Malacologia 4:1-172. map were prepared by M. Ryan (NMNH, Invertebrate TAYLOR, D. W. 1981. Freshwater mollusks of California: a Zoology); S. Escher inked anatomical drawings. Collecting distributional checklist. California Fish and Game 67:140- permits were provided by Department of Fish and Game 163. (State of California). Partial support for fieldwork and TAYLOR, D. W. 1987. Fresh-water molluscs from New Mexico laboratory studies was provided by contracts issued to the and vicinity. New Mexico Bureau of Mines and Mineral Resources Bulletin 116:1-50. author by Department of Fish and Game (State of Cali- TAYLOR, D. W. & G. R. SMITH. 1981. Pliocene molluscs and fornia), and California Department of Parks and Recre- fishes from northeastern California and northwestern Ne- ation. Assistance with fieldwork was provided by S. Ellis vada. University of Michigan, Museum of Paleontology (California Department of Fish and Game), J. Kerbavaz Contributions 25:339-413. (California Department of Parks and Recreation), and D.